Descent of the Testis
John M. Hutson • Jørgen M. Thorup
Spencer W. Beasley
Descent of the Testis
Second Edition
John M. Hutson
Royal Children’s Hospital
University of Melbourne
Parkville, Victoria
Australia
Jørgen M. Thorup
Univ. Hospital of Copenhagen
Rigshospitalet
København Denmark
Spencer W. Beasley
University of Otago Christchurch
New Zealand
The first edition of the Work was published in 1992 by HODDER ARNOLD with the following title: Descent of the Testis.
ISBN 978-3-319-25908-6
DOI 10.1007/978-3-319-25910-9
ISBN 978-3-319-25910-9 (eBook)
Library of Congress Control Number: 2015957429
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Preface
Descent of the testis, or lack thereof, is a subject of great controversy in embryology, endocrinology, and paediatric surgery. For such a common abnormality as undescended testis, it is surprising that there still remain so many unanswered questions, more than 20 years after our first edition (Table 1).
This book aims to present up-to-date evidence from our own and other laboratories of the normal mechanisms of testicular descent (Chaps. 2 and 3), along with a brief description of the evolution of descent and the early history of research into its cause (Chap. 1).
Chapters 4–10 describe the clinical problems of undescended testes and give our own, necessarily biased, opinion of the best ways to diagnose and treat the anomaly. We have not emphasised hormonal treatment, which has been vigorously proposed by others in recent years, for two major reasons. First, current evidence now suggests that direct or indirect stimulation of androgen secretion only causes relaxation of the cremaster muscle in “retractile” testes, but has little effect on truly undescended testes. Second, luteinising hormone-releasing hormone (LHRH) or its long-acting analogues have not been approved for clinical use in Australia, and one of the authors (JMH) has no direct experience of their use. Human chorionic gonadotrophin (hCG) is available, but is not in general use because of lack of efficacy and the need for a significant number of painful injections. Buserelin (LHRH analogue) is in use in some parts of Europe, but its efficacy remains very controversial.
The descriptions of the clinical diagnosis (Chap. 7) and the surgical treatment (Chap. 8) are given in detail, with extensive pictorial support, so as to give the trainee surgeon enough information to be able to learn exactly what we do. We have opted for a detailed description of orchidopexy as it is performed currently. Previous surgical techniques have been omitted deliberately because they were cruel, unnecessary, or caused testicular atrophy. There is now no place for such methods in modern paediatric surgery, where the goal is to perform day-case surgery with minimal pain or discomfort to the hapless infant or child. With this approach, orchidopexy can be performed without any painful injections or unpleasant memories.
Table1 Questions needing answers about testicular descent
Why does the testis descend?
How does the testis descend?
What causes undescended testes?
What is the best treatment for maldescent?
Does hormone therapy have a role in treatment and/or diagnosis?
Are there any new alternatives to surgical treatment?
When is the optimal time for treatment?
Are “retractile” testes normal or abnormal?
Do retractile/ascending testes need treatment?
What is the cause of infertility and increased malignancy?
Will fertility improve and the risk of cancer diminish with current modes of treatment?
The long-term results of current surgical treatment (Chap. 10) remain unknown, but we are optimistic that surgery in infancy will lead to a significant improvement in fertility and reduced risk of malignancy.
Chapter 11 concludes this short monograph with an attempt to summarise our answers to the list of questions given above. Inevitably, some of our answers will be found wanting in the future, but they will serve as a guide to future advances. Finally, we speculate on the ways that undescended testes may be treated in the future, based on our research into normal mechanisms of descent and the physiology of the testis.
There are many areas of the subject that we have not dealt with extensively; the reader should consult those books that address these areas directly. For a classic description of the human problems, the reader could do no better than read Scorer and Farrington’s 1971 treatise [1]: we have attempted unashamedly to emulate its style and approach. During the 1970s and early 1980s, the dominant paradigm was the central role of the hypothalamic-pituitary-gonadal axis which provided the rationale for hormone treatment. Several good multi-authored volumes appeared at that time, with extensive discussion of all endocrine aspects of the problem [2–7]. The role of hormones in treatment was questioned in 1986 by de Muinck Keizer-Schrama and Hazebroek in their joint thesis from Erasmus University, Rotterdam [8].
East Malvern, Australia John M. Hutson København, Denmark Jørgen M. Thorup Christchurch, New Zealand Spencer W. Beasley
References
1. Scorer CG, Farington GH. Congenital deformities of the testis and epididymis. London: Butterworths; 1971.
2. Abney TO, Keel BA, editors. The cryptorchid testis. Boca Raton: CRC Press; 1989.
3. Bierich JR, Giarola A. Cryptorchidism, proceedings of the Serono Symposia, vol. 25, Academic Press, London; 1979.
4. Fonkalsrud EW, Mengel W. The undescended testis. Chicago: Year Book Medical Publishers; 1981.
5. Hadziselimovic F, editor. Cryptorchidism, management and implications. Berlin: Springer; 1983.
6. Hadziselimovic F, Herzog B, Girard J. International symposium on pediatric and surgical andrology, cryptorchidism. Basle, Switzerland: on the occasion of the 125th anniversary of the children’s hospital, Basle; Eur J Pediatr. 1987;146(Supp1 2):S1–S68.
7. Schier F, Waldschmidt J, editors. Padiatrie aktuell 1: Maldescensus testis. Munchen: W Zukerschwerdt verlag; 1990.
8. De Muinck Keizer-Schrama SMPF, Hazebroek FWJ. The treatment of cryptorchidism why, how when? Theses, Erasmus University, Rotterdam; 1986.
Acknowledgements
This book would not have been possible without the dedicated assistance of Mrs Shirley D’Cruz. Many colleagues contributed photos or artwork and their assistance is greatly appreciated and acknowledged in the figure legends. Special thanks to Mr Bill Reid for all the line diagrams. The publishing companies are thanked for kindly giving permission to reproduce numerous figures. Finally, the authors thank Prof Spencer Beasley for his very kind permission to allow us to do a second edition of the book and Mr Sverre Klemp of Springer.
1 Evolution of Descent of the Testis and Early History of Research
1.1 Evolution of Testicular Descent
1.1.1 Why Is the Testis Descended at All?
1.1.2 Which Animals Have Descended Testes?
1.1.3 Stepwise Descent of the Testis
1.2 Early Work on Testicular Descent
1.2.1 The Role of the Gubernaculum
1.2.3
2 Transabdominal Migration of the Testis
2.1
4.1.3 Testis at the Level of External Annular Ring of the Inguinal Canal .
4.1.4 Ectopic Testes
4.1.5 The Retractile and Ascending Testis
4.1.6 The Ascending Testis
4.1.7 Acquired UDT
4.2 The Incidence of Undescended Testis (UDT)
4.3 What Causes Cryptorchidism?
5 Cryptorchidism and Associated Anomalies
5.1 Caudal Developmental Field Abnormalities
5.2 Hypospadias and the Testicular Dysgenesis Syndrome
5.3 Prune Belly Syndrome
5.4 Posterior Urethral Valve
5.5 Exomphalos and Gastroschisis
5.6 Spina Bifida
5.7 Cloacal Exstrophy
5.8 Cerebral Palsy
5.9 Testicular–Epididymal Fusion Abnormalities
5.10 Rare Fusion Anomalies
6.1 A Primary Anomaly or Secondary Effects?
6.2.3
6.3.2
6.4
7.3 Technique of Examination
7.3.1 The Appearance of the Scrotum
7.3.2 Location of the Testis
7.3.3 Description of the Level of the Testis
7.3.4 Examination of the Newborn
7.3.5 Examination of the Premature Infant
7.3.6 Mobility of the Testis
7.3.7 Cryptorchidism Associated with a Symptomatic Hernia
7.3.8 Apparent Cryptorchidism with Testicular Torsion.
7.4 Undescended Versus Retractile Testis
7.4.1 Testicular Size
7.4.2 Pain and Tightness in the Spermatic Cord
7.5 Clinical Features of Ectopic Testes
7.5.1 Definition
7.5.2 Perineal Testis
7.5.3 Pubopenile Testis
7.5.4 Femoral Testis
7.5.5 Inguino-Perineal Testis
7.5.6 Crossed Testicular Ectopia
7.5.7 Testicular Exstrophy
7.5.8 Ectopia of the Scrotum.
7.6 Impalpable Testis
7.6.1 Purpose of Investigation
7.6.2 Clinical
7.6.3 Ultrasonography
7.6.4
7.6.5
8.1.3 Anaesthesia
8.2 The Standard Orchidopexy
8.2.1 Approach to Inguinal Canal
8.2.2 Identification of Testis
8.2.3 Separation of Gubernaculum
8.2.4 The Hernia Sac
8.2.5 Freeing the Vessels and Vas
8.2.6 Placement of the Testis in Scrotum
8.2.8 Dressings and Aftercare
8.3 Unusual Findings During Orchidopexy .
8.3.1 Testicular–Epididymal Fusion Abnormalities
8.3.2 Ectopic Tissue
8.3.3 Absent Vas Deferens
8.3.4 Persistent Müllerian Duct Structures
8.4 Spermatic Cord of Inadequate Length .
8.4.1 Prentiss Manoeuvre
8.4.2 Two-Stage Orchidopexy
8.4.3 Fowler–Stephens Orchidopexy
8.4.4 Two-Stage Fowler–Stephens Orchidopexy
8.4.5 Laparoscopic Orchidopexy (Impalpable Testes)
8.4.6 Microvascular Anastomosis (Testicular Autotransplantation)
8.4.7 Testicular Biopsy
8.5 The Unilateral Impalpable Testis
8.6 Surgical Management in the Post-pubertal Child
8.7 Indications for Orchidectomy
8.8 Complications
9.2
Evolution of Descent of the Testis and Early History of Research
1.1 Evolution of Testicular Descent
1.1.1 Why Is the Testis Descended at All?
Early vertebrates had intra-abdominal gonads, regardless of sex. With subsequent evolution of sexual differences in mammals and marsupials, the testis has acquired a different position from the ovary, presumably because this different position has conferred some biological advantage. It is not known exactly why evolution has produced descended testes outside the abdominal cavity, although there is little doubt that the scrotum has become a highly specialised, low-temperature environment. Bedford [1] proposed that descent of the testis is a secondary phenomenon accompanying descent of the epididymis. Since early vertebrates, such as fish, probably ejaculated the semen into the environment, spermatozoa may have acquired advantage by being adapted to environmental temperatures below that of the body: the caudal epididymis, which stores spermatozoa, also might benefit from adaptations to cooler temperatures. In animals where the testis is not normally descended into a pendulous scrotum, the caudal epididymis is the most superficial genital organ and hence is maintained at a lower temperature than the internal organs. The testis which produces spermatozoa also may have acquired evolutionary advantage from being at a lower temperature.
Most mammals have acquired numerous sophisticated anatomical and physiological adaptations of the scrotum to keep the epididymis and testis cool (Table 1.1), many of which are present in the human. Because the human testis has inherited major adaptations fitting it to exist at a lower temperature than the body core, any disruption of normal descent would be expected to upset testicular physiology, as is discussed in Chap. 6.
The different anatomical and physiological characteristics found in different species are important for an overall understanding of human testicular descent because they reveal the various phases through which the mechanism has passed. Some of these phases may be concealed by idiosyncratic anatomical features, but the basic mechanism of testicular descent should be present in increasing complexity.
© Springer International Publishing Switzerland 2016
J.M. Hutson et al., Descent of the Testis, DOI 10.1007/978-3-319-25910-9_1
Table 1.1 The scrotum as a low-temperature environment
Character
Thin, pigmented skin
No subcutaneous fat
Absent hair/fur (e.g. bull/rat)
Pampiniform plexus
Cremaster muscle
Dartos muscle
Fat pad in inguinal canal (e.g. rat)
Fat pad between testis and epididymis (rat)
Processus vaginalis closure (primates)
Physiological role
Heat loss by conduction/radiation
Heat loss by conduction
Heat loss over caudal epididymis
Countercurrent heat exchange
Controls testicular position in response to external temperature
Controls scrotal dependency in response to external temperature
Insulates testis from abdominal cavity
Keeps testis outside abdominal cavity 1 Evolution of
Insulates caudal epididymis from testis
1.1.2 Which Animals Have Descended Testes?
Descent of the testis from its initial intra-abdominal position on the ventromedial aspect of the urogenital ridge to an external, subcutaneous scrotum occurs only in mammals. Other vertebrates, such as fish and birds (Fig. 1.1), have testes which remain inside the abdomen. Monotremes, such as the platypus and the spiny anteater or echidna (Fig. 1.2), form an intermediate group between mammals and other vertebrates and have high, intra-abdominal testes [2]. By contrast with the platypus and echidna, modern marsupials such as the kangaroo exhibit complete testicular descent to an external scrotum, albeit in the groin rather than the perineum (Fig. 1.3) [3, 4].
Some eutherian mammals have intra-abdominal testes which remain in their original position on the urogenital ridge. These include such widely different animals as the elephant and the rock hyrax (a small African herbivore) [5] (Fig. 1.4). The elephant (Fig. 1.5) and the hyrax appear to have no structure analogous to the gubernaculum. Aquatic mammals (porpoises, dolphins, and whales) have testes which are partially descended within the abdomen [6, 7] (Fig. 1.6a, b). Meek has speculated that aquatic mammals had descended testes during an earlier epoch, but this feature has been lost to a variable extent in subsequent millennia.
In the narwhale (Fig. 1.6c) [7] and the prairie dog [8] (Fig. 1.7), the testes are lateral to the bladder neck, just inside the inguinal region. Hedgehogs, moles, and shrews have testes lying lateral to the bladder; in spring and presumably in response to a surge of androgens, the enlarged testes protrude temporarily into sacs near the base of the tail [9] (Fig. 1.8). Sloths and armadillos have testes between the bladder and rectum [10].
The testes of the chinchilla are just inside the inguinal aperture, but the caudal epididymis protrudes into a thin-walled scrotal diverticulum [11 ] (Fig. 1.9 ). Some animals have testes in a subcutaneous pouch which communicates freely with the peritoneum, as is the norm for rats and mice. The southern elephant seal (Mirounga leonina Linnaeus) has two separate inguinal pouches rather than a single scrotum [ 12 ].
Fig. 1.1 The intra-abdominal gonads of the domestic fowl. (a) The large testes are high on the posterior abdominal wall of an adult rooster, near the normal position of kidneys in a mammal. Note that the testes are larger than the heart, shown at the top of the picture. (b) The single (left) ovary of the hen, showing eggs at various stages of development. The single (left) oviduct contains several eggs in transit
Even where a formed scrotum is present, its exact site and degree of development varies widely. The hyaena, for instance, has two posterior shallow scrotal pouches [13], while the kangaroo has a large prepenile scrotum in the groin located directly over the external inguinal rings.
In some mammals, the testicular position relative to the scrotum varies with sexual maturity and season. In the bear, for instance, the testes are scrotal from infancy but are held close to the body, except in the adult during the breeding season, when they are pendulous because of scrotal relaxation. The testes of the rhesus macaque descend prenatally into the scrotum but then reascend to the inguinal canal after birth, only to re-enter the scrotum finally at puberty (Fig. 1.10) [14]. This suggests that the abnormality of ‘ascending’ testes (that descend into the scrotum at puberty) in the human is normal in the macaque.
As can be seen from this brief summary, only a small number of highly specialised mammals have no testicular descent, e.g. monotremes, hyrax, elephant, and possibly some insectivores. Partial intra-abdominal descent occurs in the Cetacea (porpoises, dolphins, and whales). The testis is inguinal in position in edentates, some insectivores, and some ungulates; these animals probably have ancient origins, despite their highly specialised characteristics. Other animals have a subcutaneous pouch rather than a true scrotum and exhibit periodic or seasonal descent of the testes.
1 Evolution of Descent of the Testis and Early History of Research
Fig. 1.2 Gonadal position in the monotreme echidna. The testes and ovaries are in a similar intraabdominal position near the kidneys (Redrawn from Renfree [43]). B bladder, U ureter
Fig. 1.3 Gonadal position in the marsupial kangaroo. (a) In the male, the testis is fully descended into a prepenile scrotum, while in the female, the ovary remains near the kidney. (b) The male red kangaroo, showing the descended testes in the scrotum in the inguinal region, in front of the penis
Fig. 1.4 (a) The testicular position in the rock hyrax (Redrawn from Williams and Hutson [4]). (b) The rock hyrax, a small herbivore living in rocky outcrops in parts of Africa and one of the few mammals where the testes are in the same position as ovaries
Fig. 1.5 The intraabdominal testes of the African elephant (Reproduced with permission from Short et al. [44])
Fig. 1.6 Aquatic mammals (Cetacea), such as (a) the porpoise, (b) the fin whale, or (c) the narwhale, have testes that descended to a variable extent but are still intra-abdominal (Redrawn from Williams and Hutson [4])
Fig. 1.7 ( a ) The prairie dog has testes that have descended down to beside the bladder neck. ( b ) The prairie dog emerging from its burrow (Photo courtesy of the Royal Melbourne Zoo) 1 Evolution of Descent of the Testis and
In the large number of modern mammals, e.g. marsupials, ungulates, carnivores, and primates, the testes have descended permanently into a definite scrotum (Fig. 1.11), which in many mammals is located in the same place (i.e. just outside the inguinal canal) as the mammary glands. While life-long patency of the processus vaginalis that enables retraction of the testes back into the abdomen is normal in rodents, the processus vaginalis of apes and humans has become obliterated.
Fig. 1.8 The testis of the tree shrew is located inside the inguinal canal but can descend into a subcutaneous pouch (Redrawn from Williams and Hutson [4])
During evolution, the testes have migrated from their initial position in the embryonic urogenital ridge, which in most species is revealed in the adult by the position of the ovary. As there are a small number of mammalian species with testes in the same position as the ovary and a large number with testes in the inguinal region, it would appear that descent to the inguinal region is the first evolutionary phase of descent. It is likely that these animals evolved a mechanism for testicular descent to this point by growth of the gubernaculum [15]. Because the more-recently evolved mammals have testes outside the inguinal abdominal wall in a specialised scrotum, we can assume that further descent from the inguinal region offered further evolutionary advantage. In addition, migration of the testis beyond the inguinal region is related to migration of the gubernaculum, which is a process that is completely different from the gubernacular enlargement seen in intra-abdominal migration.
1.1.3 Stepwise Descent of the Testis
The stepwise descent of the testis during mammalian evolution parallels the stepwise descent seen in the modern mammalian fetus, such as the mouse and the human
Fig. 1.9 The chinchilla’s testis is just inside the inguinal ring, but the caudal epididymis protrudes through. There is a fat pad on the top of the epididymis, and the vas deferens joins the back of the lower urinary tract (Redrawn from Williams and Hutson [4])
[16]. Between about weeks l 0 and 15 of gestation in the human, the testis remains close to the future internal inguinal ring because the enlarging gubernaculum in the male acts as an anchor. On the other hand, the gubernaculum in the female fetus remains thin and tenuous, and the ovary (in most species but not the human) shifts away from the inguinal region as the fetus enlarges [17]. The elongation of the thin gubernaculum to form a round ligament in the female is the reverse of the broad, short gubernaculum in the male that remains at a fixed length. Around 25–28 weeks gestation, the testis descends rapidly through the inguinal canal and then migrates more slowly from the external inguinal ring to the scrotum, arriving there at 35–40 weeks. This phase of descent is preceded by migration of the gubernaculum from the inguinal canal to the scrotum.
The stepwise descent of the testis observed in human embryos and laboratory animals led to the concept that testicular descent may occur in two or more phases. This concept was proposed initially by Wensing [ 15 , 18 ] and one of the
Fig. 1.10 (a) In the macaque, the testis descends into a scrotum at birth but then ascends to the groin until puberty, when permanent descent occurs. This animal shows that ‘ascending testis’ with final descent at puberty can be normal, as compared to the human where it is abnormal (Reproduced with permission from Ref. [14]). (b) An albino adult male macaque, showing the scrotal position of the testes (Photo courtesy of the Royal Melbourne Zoo)
authors [ 19 , 20 ]. Wensing’s observations of the enlargement (or ‘swelling reaction’) of the male gubernaculum in the pig fetus and other species, coupled with observations of rodents by Habenicht and Neumann [18 ], suggested that testicular descent was not only stepwise morphologically but also stepwise in its hormonal control. The naturally occurring mutation in the androgen receptor gene in the testicular feminising (TFM) mouse, in which the androgen receptor is completely non-functional, provided crucial evidence for stepwise testicular descent because no tissues in this mouse are able to respond to circulating androgens [ 19 ]. In this animal the swelling reaction of the gubernaculum occurs relatively normally, and the testis resides near the inguinal region beside the bladder neck, but no migration outside the abdomen occurs. The explanation was that androgens are required for migration from the groin to the scrotum, but that other hormonal factors control intra-abdominal migration and enlargement of the gubernaculum [ 19 ].
The detailed evidence for multiphasic testicular descent, both morphological and hormonal, is discussed in Chaps. 2 and 3: here we summarise some of the historical evidence which has been accumulated over the last two centuries [3, 4].
Fig. 1.11 The testis in ungulates descends into a pendulous scrotum before birth. (a) In this prize bull, the hairless scrotum is quite conspicuous and is located in the same anatomical site as the mammary glands in a cow (b)
1.2 Early Work on Testicular Descent
1.2.1
The Role of the Gubernaculum
The study of testicular descent began in earnest with the observations of John Hunter [21] (Fig. 1.12). In 1762 and 1786, he described the fetal testis and epididymis in the abdomen and provided the first accurate description of the gubernaculum. He found that the fetal testis was connected to the abdominal wall by a ligament, or so-called gubernaculum testis, because it appeared to direct the course of the testis during descent. Hunter thought that the gubernaculum was vascular and fibrous and covered by fibres of the cremaster muscle.
1.2 Early Work on Testicular Descent
Fig. 1.12 John Hunter, Scottish surgeon–scientist, who first described the gubernaculum. A close up from an engraving by G.H. Adcock, based on a painting by Sir Joshua Reynolds
The relationship between the gubernaculum and the developing cremaster muscle within it was the subject of much controversy, in part related to anatomical differences between species. Seiber [22] described ascending muscular fibres in the gubernaculum by extrapolation from dissections of adult non-human mammals. Curling [23] described the gubernaculum as a soft, solid cone which varies in shape and size at different stages of descent. He found a bulky central part composed of primitive cellular tissue (i.e. embryonic mesenchyme) surrounded by a layer of muscular fibres and another layer of ‘mesenchyme’, the whole being invested by peritoneum except posteriorly. Cloquet and Carus [24, 25] believed that the cremaster was formed mechanically from the passage of the testis through the abdominal wall, which pulled loops of muscle fibres off the conjoint tendon made up of the internal oblique and transversus abdominis muscles.
Migration of the gubernaculum from the inguinal region to the scrotum was well recognised in the nineteenth century. Cleland [26] dissected human fetuses of 5 and 6 months’ gestation and found that the fetal gubernaculum ended in the inguinal abdominal wall and did not extend directly from the testis to the scrotum. Hence, the alternative name for the gubernaculum became the genito-inguinal ligament. Cleland believed that the gubernaculum migrated ahead of the testis into the scrotum, creating a space lined by peritoneum. He did not think that the cremaster muscle pulled the testis into the scrotum. (It is not surprising that our current
understanding of the gubernaculum, as a structure migrating to the scrotum with the testis inside it in the processus vaginalis, is effectively the same as that described by Cleland more than 150 years ago.)
One theory which was popularised by Lockwood [27] was that differential growth of the posterior abdominal wall relative to the pelvis meant that the testis remained stationary while its surroundings grew further apart. Lockwood did not believe that the gubernaculum contracted but imagined that the gubernaculum passively held the testis near the inguinal region. In subsequent years, this passive effect of relative growth was interpreted to mean that this phase of descent was unimportant. However, anchoring of the gonad by the gubernaculum fails to occur in the female fetus, indicating that this effect in the male is hormonally mediated. In fact, in the female fetus, the ovary actually ascends relative to the position it occupied initially in early development.
Lockwood also speculated that the gubernacular muscle must serve some purpose, leading to the idea that the different final location to the testis may be caused by a fan-shaped gubernaculum. Although he was unable to demonstrate these socalled Tails of Lockwood on dissection, he postulated their existence as an explanation for the observation that the gubernaculum becomes attached in a range of different positions [28, 29]. Now that it is known that the gubernaculum actually migrates to the scrotum, the ‘Tails of Lockwood’ should no longer be considered a valid concept [29], and the mechanical phases of testicular descent are better understood (Fig. 1.13).
Coley [30] and Sebileau [31] did not believe that an ectopic position of the testis was caused by an aberrant gubernaculum. Many surgeons and authors believe that an undescended testis or ectopic testis is caused by mechanical obstruction [32–34].
1.2.2 The Role of Hormones
In the early twentieth century, the advent of hormonal studies of sexual development undermined the credibility of previously held mechanical and anatomical concepts [35, 36]. In an effort to ‘explain’ testicular descent in endocrinological terms, these earlier mechanical hypotheses were discarded. Initially, the simplest hormonal concept was that male androgens under pituitary control simply caused testicular descent [37]. This led to attempts to treat boys with undescended testis with crude preparations of human chorionic gonadotrophin (hCG) and, once these steroids had been synthesised, with androgens. Early optimistic reports of success of these treatments were eventually replaced with more realistic and consistently disappointing reports.
1.2.3 Phasic Control
Experimental evidence from the fetus failed to support the idea that testicular descent was controlled by testicular androgens alone [15, 18, 38, 39]. Such
Fig. 1.13 The mechanical steps of testicular descent. Schema showing the testicular position and gubernaculum at different stages of fetal development. (a) At 10 weeks the testis is in the urogenital ridge with both Wolffian and Müllerian ducts, and the gubernaculum (genito-inguinal ligament) attaches the testis to the abdominal wall. (b) By 15 weeks the Müllerian duct has regressed, and Wolffian duct is developing into an epididymis and vas deferens. The gubernaculum has enlarged, which holds the testis near the future inguinal canal. The processus is beginning to grow into the gubernaculum. (c) At 30 weeks the gubernaculum is migrating to the scrotum with the testis inside the processus vaginalis, which is elongating within the gubernaculum so that the intra-abdominal testis can reach the scrotum while still inside the processus vaginalis. (d) By 35 weeks migration is usually complete, and the gelatinous gubernaculum has resorbed, leaving the processus vaginalis, which becomes attached to the scrotal skin. The proximal processus obliterates, leaving the testis in a satellite peritoneal cavity (tunica vaginalis) in the hemiscrotum
discrepancies eventually led to the proposal of multihormonal control [40]. Evidence from our own laboratory then suggested that testicular descent occurred in two separate phases, each of which was controlled by different hormones [41], and this is now widely accepted [42] and described in Chaps. 2 and 3
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G/23905 Tovey, E. H.
G/3298 Towler, B
G/4559 Town, A C
G/4932 Town, J
L/7944 Town, Y T E
T/20369 Townsend, T
G/22061 Toyer, E C J
G/29238 Tranter, E
G/25226 Treadgold, A.
T/203770 Tredwell, G. A.
G/1778 Tressidder, J. A.
T/242015 Trew, E. C., M.M.
G/26600 Trewin, F. S.
G/12938 Trewin, H. C.
G/20053 Trice, A. H.
T/242960 Trice, L.
G/139 Trice, W.
G/20174 Triggs, F
G/7914 Triptree, A G
G/2302 Tritton, C J
G/409 Trott, H J
G/2511 Trowbridge, C H
G/12567 Trubody, P
G/410 Trull, A W H
T/3009 Trumpeter, A. J.
G/657 Tucker, A. H.
T/203479 Tucker, J. S.
L/6965 Tucker, J W
L/8090 Tucker, R G
T/204014 Tuffley, F
T/202063 Tuffrey, H
G/15699 Tugwell, W T B
G/4731 Tully, F
G/6741 Tunnicliffe, A
S/429 Tunnicliffe, W.
T/241000 Tunstall, H. W.
G/577 Tupp, C. J.
G/23902 Tupper, A.
G/389 Tupper, C. E.
T/241532 Turner, D. J.
G/19042 Turner, E.
L/10406 Turner, G.
G/18930 Turner, G.
G/8377 Turner, G. S.
G/4585 Turner, H W
G/7959 Turner, J
G/13071 Turner, J
G/10629 Turner, J
G/8249 Turner, M J
G/2516 Turner, R H
G/7219 Turner, R
G/17717 Turner, S.
G/5600 Turner, S. E.
G/3864 Turner, W. A.
G/21064 Turner, W. M.
G/14086 Turney, M.
L/9588 Turrell, J.
G/29236 Turton, G.
G/23904 Tutin, T. A.
G/5495 Tutt, F.
S/10403 Tutt, T
G/19051 Twaite, G
G/22099 Tween, W
G/3973 Twigg, T W
G/2313 Twin, C F
G/4878 Twinn, W C
G/9329 Twyman, C H
G/5140 Twyman, G. H.
G/5070 Twyman, P. C.
G/4651 Twyman, T.
G/21932 Tyler, A
G/8682 Tyler, A C
G/13176 Tyler, A H
G/15525 Tyson, W
T/3380 Uden, A
G/9893 Uden, E G
T/240388 Underdown, G
G/15807 Underhill, C
T/24284 Upshall, C E
G/19004 Upson, H
G/1760 Upton, A. G.
G/9118 Usherwood, H. C.
T/1580 Van Rooyen, G. J. C.
G/6662 Vaughan, F. C.
G/4962 Veitch, H. J.
T/4461 Velvick, C. E.
G/1380 Veness, J.
T/203965 Venn, M.
L/9181 Venton, F.
G/2128 Vernon, R. A. W.
L/10527 Vicary, L. J.
T/1746 Vidler, J. A.
G/15702 Vidler, S C
G/24588 Viggor, F
G/10426 Vinall, E T
L/8897 Vinall, G
G/831 Vincent, A J
L/10303 Vincent, F W
G/18880 Vincent, S G
T/204552 Viner, A.
G/13089 Vining, C. H. E.
G/14123 Virgin, F. T.
G/21009 Voller, G.
T/240808 Vousden, C. R.
G/5795 Vousden, H.
G/645 Vousden, R. F.
G/8550 Vousden, W. H.
G/3503 Wade, G. T.
G/1680 Wade, W.
G/8200 Waghorn, F.
L/7239 Waghorn, J E
G/11443 Waghorn, T E
G/5967 Waight, J H
T/200855 Wakelin, F
G/14375 Waldie, J D
G/3521 Walker, A J
G/3397 Walker, B
T/241300 Walker, J. H.
G/14868 Walker, J. H. A.
L/8766 Walker, R. J.
G/2870 Walker, R.
S/98 Walker, T.
G/20967 Walker, T. G.
L/7233 Walkom, G. A.
G/4144 Wallace, D.
L/7865 Wallace, W. J.
G/14651 Waller, J. R.
G/20800 Wallis, A
G/1296 Wallis, H W
G/11693 Wallis, W A
S/543 Walter, E
G/11667 Walter, R
G/13284 Walton, J
G/7626 Wanstall, T F
G/18902 Want, J.
G/10875 Warby, W. R.
G/24707 Ward, A. E.
L/10094 Ward, C.
T/206060 Ward, F.
G/8156 Ward, G.
G/17660 Ward, J. H.
T/241539 Ward, J. W.
T/203220 Ward, S. G.
L/7880 Ward, T C S
L/8624 Ward, W J R
L/8571 Warden, C H
G/29253 Wardle, J W
G/11460 Ware, A
G/35685 Ware, J G
G/1611 Waring, S
G/13629 Warman, T.
G/13431 Warman, W. R. H.
G/1007 Warne, R. G.
G/637 Warren, P
G/1015 Warry, T V
G/20035 Warwick, F H
G/14922 Warwick, J W
G/1616 Washbrook, A
G/11668 Wassell, H J
G/5792 Watches, P
G/29240 Waterall, T. W.
T/270967 Waterhouse, W. W.
G/1192 Waters, H.
G/18888 Waters, R. J.
T/270439 Waters, T. J.
G/4317 Watkins, H.
G/798 Watkins, H. J.
S/229 Watson, B.
G/190 Watson, F.
G/171 Watson, G.
G/1702 Watson, H E
T/201021 Watson, J
G/6429 Watson, J
G/7007 Watson, J
G/6190 Watson, J H
G/18662 Watson, R C
G/2044 Watson, W
G/1932 Watson, W. H. J.
G/13257 Watts, A.
G/22619 Watts, F. W.
L/7886 Watts, R. W.
G/6289 Watts, S. A.
G/5428 Watts, W. E. A.
T/2181 Wayte, J.
G/15753 Weare, L. T.
G/9510 Weatherall, A. P.
T/242824 Weatherill, E
G/18206 Weaver, A G
G/13599 Weaver, R
G/12666 Weavers, T J
G/11957 Webb, A A
G/4723 Webb, E C
G/4616 Webb, E E
G/14015 Webb, E. S.
